The Effects of Drought and Shade on the Performance,Morphology and Physiology of Ghanaian Tree Species

In tropical forests light and water availability are the most important factors for seedling growth and survival but an increasing frequency of drought may affect tree regeneration. One central question is whether drought and shade have interactive effects on seedling growth and survival. Here, we present results of a greenhouse experiment, in which seedlings of 10 Ghanaian tree species were exposed to combinations of strong seasonal drought (continuous watering versus withholding water for nine weeks) and shade (5% irradiance versus 20% irradiance). We evaluated the effects of drought and shade on seedling survival and growth and plasticity of 11 underlying traits related to biomass allocation, morphology and physiology. Seedling survival under dry conditions was higher in shade than in high light, thus providing support for the “facilitation hypothesis” that shade enhances plant performance through improved microclimatic conditions, and rejecting the trade-off hypothesis that drought should have stronger impact in shade because of reduced root investment. Shaded plants had low biomass fraction in roots, in line with the trade-off hypothesis, but they compensated for this with a higher specific root length (i.e., root length per unit root mass), resulting in a similar root length per plant mass and, hence, similar water uptake capacity as high-light plants. The majority (60%) of traits studied responded independently to drought and shade, indicating that within species shade- and drought tolerances are not in trade-off, but largely uncoupled. When individual species responses were analysed, then for most of the traits only one to three species showed significant interactive effects between drought and shade. The uncoupled response of most species to drought and shade should provide ample opportunity for niche differentiation and species coexistence under a range of water and light conditions. Overall our greenhouse results suggest that, in the absence of root competition shaded tropical forest tree seedlings may be able to survive prolonged drought.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

森林植物耐荫性及其形态和生理适应性研究进展

耐荫性是植物在低光环境下的生存和生长能力,对森林植物群落演替起重要作用,植物对遮荫的适应机制已成为生态学的研究热点.本文综述了森林植物的耐荫性及其在形态和生理方面的适应性,分析了森林植物在生长性状、生物量分配、树冠结构、叶片形态生理、叶片解剖结构、光合参数、碳水化合物分配、水分和养分的利用等方面对遮荫产生的可塑性响应,最后对目前研究存在的问题进行了分析,展望了未来的研究内容和方向.     

Influence of light level on the growth and morphology of saplings in a panamanian forest

Leaf spacing and aboveground growth were monitored in saplings of ten species in a range of light environments in a Panamanian lowland humid forest. One- to 2-m-tall individuals of the chosen species had intermediate to large leaves on stems with few or no branches. Saplings in high light environments grew faster in height and produced more biomass per unit leaf area than shaded saplings for all species. These growth responses involved morphological plasticity with greater extension per unit biomass increment increasing the height growth rate of gap-grown saplings and greater biomass allocation to leaves decreasing the whole plant light compensation point in shade. The relative performance of the species also varied across the light gradient and was related to differences in leaf lifespan and specific leaf mass. Light-demanding species grew as rapidly in shade as shade-tolerant species, but the shorter leaf lifespan of the former necessitates higher production rates to maintain a given leaf area, largely excluding light-demanders from shaded understory locations. Height growth rate was positively correlated with leaf spacing for each species, and differences between species in the height growth rate-internode length relationship were related to interspecific differences in specific leaf mass. Thus, sapling growth histories may be inferred from their morphologies.     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size

An evergreen oak species, Cyclobalanopsis multinervis, and a deciduous oak species, Quercus aliena var. acuteserrata were grown from acorns under two light levels (full sunlight and shade at about 18 % of full sunlight, simulating the light intensities in forest clearings and gaps, respectively) for one growing season. Three hypotheses were tested: (i) the deciduous species grows faster than the evergreen species in forest gaps and clearings; (ii) the deciduous species responds more strongly in terms of growth and morphology to variation in light climate than the evergreen species; and (iii) seedling size is positively correlated to acorn size. The results showed: (i) at both light levels, the deciduous seedlings gained significantly more growth in biomass and height than the evergreen seedlings; (ii) both species produced significantly more biomass in full sunlight than in shade, without showing any significant difference in height between treatments. Increase in light intensity improved the growth of the deciduous seedlings more strongly; (iii) at a similar age, the deciduous seedlings showed a greater response in leaf morphology and biomass allocation to variation in light levels, but when compared at a similar size, biomass allocation patterns did not differ significantly between species; (iv) bigger acorns tended to produce larger seedlings, larger leaf sizes and more leaf area, between and within species. These differences demonstrate that the deciduous species is gap-dependent and has the advantage over the evergreen species in forest gaps and clearings.     

Morphological and biomass characteristic acclimation of trident maple (Acer buergerianum Miq.) in response to light and water stress

Acer buergerianum Miq. (Trident maple) is a native species of China with a large distribution, but exist in small population. Water and light are two important factors limiting plant growth and are crucial in the framework of forest regeneration. However, there is no consensus on how shade interacts with drought. Four hypotheses in the recent literature variously predict that shade will have a stronger, weaker or equal impact on seedlings under drought stress. This study investigated the interactive responses of A. buergerianum to light and water focusing on seedling growth, leaf morphology and biomass partitioning by performing a growth experiment in pots with different water supply regimes [15, 35, 55, 75, 95 % of field capacity (FC)] combined with two light regimes (10 and 66 % of full sunlight). After 123 days treatment, the results showed that shade greatly reduced growth and biomass, in contrast enhancing the amount of chlorophyll, the amount of water in the leaves, and the specific leaf area. Drought reduced growth, biomass, and the bulk of the leaves. Most leaf traits and biomass characteristics had strong interactions in their responses to light and water treatments. Allometric analysis revealed that water and light had no effects on root to shoot ratios, main root to lateral root ratios, and root mass ratios. Shade alleviated the negative impact of drought. A. buergerianum successfully adapted to the various light and water conditions. We recommend a water supply above 15 % FC to keep the seedlings vigorous, under both sunlight conditions.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.     

Variation for phenotypic plasticity among populations of an invasive exotic grass

Phenotypic plasticity is a common feature of plant invaders, but little is known about variation in plasticity among invading populations. Variation in plasticity of ecologically important traits could facilitate the evolution of greater plasticity and invasiveness. We examined plasticity among invasive populations of Microstegium vimineum (Japanese stiltgrass), a widespread and often dominant grass of forests in the eastern U.S. with two separate experiments. First, we exposed seven Microstegium populations to a drought treatment in growth chambers and monitored growth and physiological responses. Then, we established a greenhouse experiment using a subset of the populations; two that exhibited the most divergent responses and one intermediate population. In the greenhouse, we manipulated drought and shade and evaluated biomass production and specific leaf area (SLA). Microstegium exhibited plasticity for biomass production and SLA in the greenhouse experiment, and populations significantly varied in the degree of plasticity under drought and shade treatments. Two populations significantly increased biomass production under favorable conditions, unlike the third population. The most productive populations also responded to shade stress via greater SLA, possibly allowing for greater utilization of available light, while the third population did not. These results show that Microstegium can exhibit plastic responses to environmental conditions. Moreover, variation for plasticity among populations provides the potential for further evolution of plasticity. Future studies should focus on the relative importance of plasticity for the success of Microstegium and other plant invaders and evaluate post-introduction evolution of plasticity.     

不同光环境对麻栎和刺槐幼苗生长和光合特征的影响

通过搭建遮阳棚设置3个光照强度,模拟森林幼苗生长的林缘、林窗和林下光环境,研究麻栎和刺槐幼苗形态结构、光合特性、生物量积累及其分配对不同光照强度的响应。结果表明,极度弱光环境限制了幼苗的株高、基径、总叶面积、冠面积、叶面积指数和总叶数等形态指标的增长,与全光环境相比,适度遮荫有利于幼苗的形态生长,光照对麻栎形态的异速生长曲线影响较小,刺槐较麻栎的曲线变化更明显一些,弱光降低了异速生长指数。随遮荫程度的增加,麻栎的净光合速率、蒸腾速率、气孔导度逐渐降低,刺槐的净光合速率则在适度遮荫下最大,两者的光能利用效率均显著增大,水分利用效率先升后降。随光照强度的减弱,麻栎各器官的生物量积累降低,叶面积比率的增加和根冠比的减少有利于增加光能的捕获,而刺槐的生物量积累和分配则在适度遮荫下达到最优;各叶绿素含量均显著上升,而叶绿素a/b则逐渐下降。麻栎和刺槐显示出不同的生存策略和光利用策略,在一定程度上可以揭示它们在阔叶林群落中不同的演替地位。     

攀援植物绞股蓝幼苗对光照强度的形态和生长反应

 利用遮阳网产生光照强度梯度,以研究攀援植物绞股蓝(Gynostemma pentaphyllum)幼苗对光梯度的形态和生长反应。结果表明：1)相对生长速率、净同化速率和总生物量随光照强度减弱而降低,总叶面积、比茎长、株高、分枝角度、叶面积率和比叶面积却增加;2)株高生长与相对生长速率成负相关;3)幼苗生物量分配对光梯度的反应不敏感。这些结果意味着绞股蓝幼苗的形态和生长反应对不同光环境具有可塑性,比茎长和株高随光照减弱而增加有利于绞股蓝幼苗“寻找”到外界支持物。     

光照和氮素对喜旱莲子草形态特征和生物量分配的影响

研究了两个光照梯度和3个土壤氮素水平交互作用对喜旱莲子草(Alternan thera philoxeroides(Mart.)Griseb.)形态特征和生物量分配的影响。结果表明,全光照促进喜旱莲子草总生物量的积累,但在遮荫条件下,喜旱莲子草可以通过增加株高、光合叶面积和改变生物量分配来适应弱光生境。土壤中氮素含量对喜旱莲子草生长有明显影响,总生物量、株高、叶面积、茎生物量比和叶生物量比等随土壤氮素水平增加而增加。光照和氮素的交互作用对总生物量、根生物量比、茎生物量比和叶生物量比也有显著影响。随着氮素水平的增高,遮荫和高光照处理下喜旱莲子草的叶面积、总生物量和叶生物量比之间的差异减小,而株高和根生物量比之间的差异增大。此外,光照强度对茎生物量比的影响具有明显的氮素浓度依赖性,低氮条件下,茎生物量比在高光照处理下显著高于遮荫处理,而在中氮条件下,遮荫处理却显著高于高光照处理,且在高氮处理下其差异进一步加大。这些结果表明喜旱莲子草在高氮素环境下能够通过形态可塑性和生物量分配模式的改变来适应弱光环境所带来的不利影响。研究结果不但可为研究喜旱莲子草对异质生境的入侵机制提供资料,也可为进一步研究喜旱莲子草的入侵和扩散与农业等生态系统中土壤氮素残留的关系提供参考。       

A morphophysiological analysis of the effects of drought and shade on <Emphasis Type="Italic">Catalpa bungei</Emphasis> plantlets

The interactive effects of shade and drought on the morphological and physiological traits of Catalpa bungei plantlets were assessed. Seedling growth, biomass, biomass allocation, leaf morphology, chlorophyll (Chl) content and gas-exchange parameters were measured in plants raised for 3 months under three light levels [80% (HI), 50% (MI), 30% (LI)] and two water levels [moisture (M) and drought (D)]. The results showed that shade greatly decreased growth, biomass, leaf area (LA) and Chl a/b; increased specific leaf area (SLA) and Chl content; and reduced photosynthetic rate (P _n). Drought reduced the growth, biomass, LA, SLA, Chl a/b, P _n, stomatal conductance (G _s), transpiration rate (T _r) and intercellular carbon dioxide concentration (C _i) and increased the Chl content. Stomatal closure was an early physiological response to water stress. Light, water and their interaction significantly affected plant traits and their bivariate relationships. The phenotypic plasticity index of light (0.47) was much higher than that of water (0.21), indicating that light was the main driver of the variations observed. Under drought stress, growth, biomass, leaf and stem biomass allocation significantly decreased in the HI and MI environments, whereas no significant difference was observed in growth or biomass parameters under the LI condition. Furthermore, no significant difference was observed in P _n, G _s, or T _r under the LI condition under water stress. Our results showed that shade did not alter the negative effects caused by drought stress in MI but did alleviate the negative effects of the LI condition. In summary, the effect of drought on C. bungei plantlets depends on the irradiance conditions.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Consequences of phenotypic plasticity vs. interspecific differences in leaf and root traits for acquisition of aboveground and belowground resources

Trade-offs between acquisition capacities for aboveground and belowground resources were investigated by studying the phenotypic plasticity of leaf and root traits in response to different irradiance levels at low nutrient supply. Two congeneric grasses with contrasting light requirements, Dactylis glomerata and D. polygama, were used. The aim was to analyze phenotypic covariation in components of leaf area and root length in response to above- and belowground resource limitation and the consequences of this variation for resource acquisition and plant growth. At intermediate shading (30 and 20% of full sunlight) the plants were able to maintain their total root length, despite a strongly increased total leaf area and a reduced biomass allocation to roots. This was associated with an unaltered or slightly increased nutrient uptake and growth. At 5.5% relative irradiance, growth was severely reduced, especially in the shade-tolerant D. polygama. The results show that constraints on acquisition capacities for aboveground and belowground resources, caused by biomass allocation, may be alleviated by plasticity in other traits such as tissue-mass density and thickness of roots and leaves. The results also suggest different adaptive constraints for phenotypic plasticity and for genetically determined interspecific variation. Phenotypic plasticity tends to maximize resource acquisition and growth rate in the short term, whereas the higher tissue-mass density and the longer leaf life-span of shade-tolerant species indicate reduced loss rates as a more advantageous species-specific adaptation to shade in the long term.     

Survival of three cool-temperate tree species in shade with lime and NPK applications

Knowledge of the site specificity of plastic reactions and shade tolerance of tree species within the complexity of natural forests is limited but critical to understanding forest regeneration. Paper birch (Betula papyrifera), yellow birch (B. alleghaniensis), and eastern hemlock (Tsuga canadensis) were planted in a randomized block design under natural deciduous canopy and within small canopy gaps with a single application of lime and NPK at time of planting in 2003. Saplings were excavated in 2004 and 2008 for analysis of biomass allocation and testing of foliar N. In 2008, mean survival rate differed significantly among species at 0.14 for paper birch, 0.57 for yellow birch, and 0.76 for eastern hemlock, respectively. Yellow birch emerged as a species of intermediate shade tolerance associated with a sharp decline in shoot development after two growing seasons, high leaf allocation of biomass, and foliar plasticity. Foliation and physiological adaptation of leaves appeared N-limited in yellow birch. Biomass allocation analysis revealed strong allometric growth in yellow birch and eastern hemlock across experimental treatments imposed. Increased soil reaction from pH 4.7 to 6.3 following lime applications had no discernable effect on the juvenile growth and survival of the three species during six growing seasons. The experiment underlined the need for experimental testing of species under natural forest canopy. Cyclical cutting of forests with moderate canopy opening is recommended for sustainable management of yellow birch and eastern hemlock.     

Do the costs and benefits of fungal endophyte symbiosis vary with light availability?

? Here, we examined whether fungal endophytes modulated host plant responses to light availability. First, we conducted a literature review to evaluate whether natural frequencies of endophyte symbiosis in grasses from shaded habitats were higher than frequencies in grasses occupying more diverse light environments. Then, in a glasshouse experiment, we assessed how four levels of light and the presence of endophyte symbioses affected the growth of six grass species. ? In our literature survey, endophytes were more commonly present in grasses restricted to shaded habitats than in grasses from diverse light environments. ? In the glasshouse, endophyte symbioses did not mediate plant growth in response to light availability. However, in the host grass, Agrostis perennans, symbiotic plants produced 53% more inflorescences than nonsymbiotic plants at the highest level of shade. In addition, under high shade, symbiotic Poa autumnalis invested more in specific leaf area than symbiont-free plants. Finally, shade increased the density of the endophyte in leaf tissues across all six grass species. ? Our results highlight the potential for symbiosis to alter the plasticity of host physiological traits, demonstrate a novel benefit of endophyte symbiosis under shade stress for one host species, and show a positive association between shade-restricted grass species and fungal endophytes.     

Growth and leaf production in the tropical palm Euterpe edulis: Light conditions versus developmental constraints

We studied developmental and environmental constraints on leaf dynamics, morphology and physiology in the monopodial tropical palm of the Atlantic Forest biome, Euterpe edulis. Plastic responses to light environments in terms of photosynthesis, leaf size, leaf life span, patterns of biomass allocation and growth were analysed. Plants were grown during 14 months in a shade house under four different growth irradiances. Plants of Euterpe edulis were able to adjust leaf demography and biomass allocation in the different light treatments. Leaf life span increased by 100 days with decreasing light levels while the rate of leaf production decreased, consistent with lower electron transport rates. At low light levels, adjustments in biomass allocation to leaf components allowed E. edulis to reduce self-shading and increase light interception. At high light plants allocated more biomass to roots, and the plants exhibited small leaf sizes when leaves were compared using an explicit ontogenetic analysis. Ontogeny constrained the maximum size that each consecutive leaf could achieve, while growth irradiance determined the rate of leaf production and other leaf traits. Consequently, there were both, developmental constraints and environmental determinants influencing leaf demography and morphology in E. edulis. The findings of this ecophysiological and demographic study are relevant to palms growing under natural conditions and help to explain the success of E. edulis in the forest understory and its absence from large gap openings. Our results not only confirm that E. edulis is a shade tolerant species, but also show that palms are able to acclimate to different growing condition as well as trees.     

Is hemiepiphytism an adaptation to high irradiance? Testing seedling responses to light levels and drought in hemiepiphytic and non‐hemiepiphytic Ficus

The epiphytic growth habit in many Ficus species during their juvenile stages has commonly been hypothesized to be an adaptation for avoiding deep shade in the forest understory, but this has never been tested experimentally. We examined growth and ecophysiology in seedlings of three hemiepiphytic (Hs) and three non‐hemiepiphytic (NHs) Ficus species grown under different irradiance levels. Both Hs and NHs exhibited characteristics of high light requiring species, such as high plasticity to growth irradiance and relatively high maximum photosynthetic assimilation rates. Diurnal measurements of leaf gas exchange showed that Hs have much shorter active photosynthetic periods than NHs; moreover, leaves of Hs have lower xylem hydraulic conductivity but stronger drought tolerance as indicated by much lower rates of leaf diebacks during the drought treatment. Seedlings of NHs had 3.3‐ and 13.3‐fold greater height and biomass than those of Hs species after growing in the nursery for 5 months, indicating a trade‐off between growth and drought tolerance due to the conflicting requirements for xylem conductivity and cavitation resistance. This study does not support the shade‐avoidance hypothesis; rather, it suggests that the canopy regeneration in Hs is an adaptation to avoid alternative terrestrial growth‐related risks imposed to tiny Ficus seedlings. The NHs with terrestrial regeneration reduce these risks by having an initial burst of growth to rapidly gain relatively large seedling sizes, while in Hs seedlings more conservative water use and greater drought tolerance for surviving the canopy environment are intrinsically associated with slow growth.     

紫茎泽兰和飞机草的形态和光合特性对磷营养的响应

比较研究了紫茎泽兰和飞机草的生长、形态、生物量分配和光合特性对磷营养的可塑性反应,及与其入侵性的关系.结果表明,两种入侵植物对磷营养变化表现出很强的可塑性和适应性.低磷时,两种植物的根生物量比增大,利于养分吸收;高磷时,两种植物的比叶面积、最大净光合速率、光饱和点、单位面积的叶绿素和类胡萝卜素含量增大,同化面积和同化能力增强,利于碳积累.相比之下,紫茎泽兰对磷的适应性更强.随供磷量的增加,紫茎泽兰和飞机草的相对生长速率、总生物量、株高、分枝数、叶面积指数和最大净光合速率均显著增大,过量磷素对上述参数抑制较小,表明两种入侵植物偏好较高的磷环境,土壤磷含量升高有利于其入侵,并在高磷时,通过增大株高、分枝数和叶面积指数荫蔽排挤本地种.在本地种基本停止生长的干季,紫茎泽兰和飞机草仍维持较高的相对生长速率.这也与其入侵性密切相关.